Long‐term body pigmentation changes on a manta ray (Mobulidae)

Manta rays (genus Manta) are listed as vulnerable on the IUCN Red List based partly on population studies that use body coloration and ventral spot patterns to identify and count individuals, as a result of such patterns being considered unique and permanent during their lifetime. The present study reports the first evidence of long‐term coloration changes of manta rays based on observations of one captive individual. Darkening skin pigmentation was documented on the side of the head, the inner side of the cephalic lobes and the pectoral fin margin of the ventral side, and spot markings appeared on the gill slits within 9 months. These changes were most likely ontogenetic or were connected to maturation. The described specimen's coloration did not fulfill the taxonomic classification criteria of either manta ray species and rather resembled two different manta ray species at the beginning and end of the study. These results show that coloration patterns of manta rays are not as stable as has been assumed previously. Further studies are needed to identify the extent of such changes for accurate identification and classification of manta rays. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 406–414.     

Running the Gauntlet: Regional Movement Patterns of Manta alfredi through a Complex of Parks and Fisheries

Manta rays (Genus Manta) are economically important for fisheries and tourism in Indonesia. These species have been listed by the International Union for the Conservation of Nature Red List as Vulnerable to extinction; therefore, human exploitation of manta rays must be regulated. A better understanding of the habitat use and movement patterns of manta rays in Indonesia is needed in order to employ effective conservation measures. To gain better insight into the movements of Manta alfredi we used ‘Manta Matcher’, an online database with an integrated automated matching algorithm, to compare photographs from 2,604 encounters of M. alfredi collected by recreational divers and dive operators throughout Indonesia over a nine-year period. This photographic comparison revealed that manta rays migrated between regional sanctuaries such as Nusa Penida, the Gili Islands, and the Komodo National Park (up to 450 km straight-line distance). The areas between these sanctuaries are heavily fished and trafficked by ships, and when manta rays travel through these regions they risk being fished and injured by ship strikes. These long-range manta ray movements suggest connectivity between M. alfredi populations in neighboring islands and raise concerns about the future management of regional populations. It is recommended that a national conservation strategy be developed to protect the remaining populations in the country.     

Seasonal occurrences of <Emphasis Type="Italic">Manta birostris</Emphasis> (Chondrichthyes: Mobulidae) in southeastern Brazil

An analysis of 79 underwater photographs of Manta birostris gathered over a period of nine years in a marine protected area in southeastern Brazil suggests a high predictability of manta ray occurrences in the region during the austral winter (June–September). The reasons for this are probably related to the seasonal oceanographic conditions, as characterized by the presence of a coastal front at the study site in winter and consequent plankton enrichment, which provides a feeding opportunity for manta rays. In addition, a melanistic individual in the Atlantic Ocean that is similar in color to the Pacific Ocean’s “black manta” is reported for the first time.     

Population estimates of Alfred mantas (<Emphasis Type="Italic">Manta alfredi</Emphasis>) in central Maldives atolls: North Male,Ari and Baa

The Alfred manta, Manta alfredi, is economically important in the Maldives due to the large number of tourists attracted to SCUBA dive and snorkel with them. It is important to understand the size of populations around main diving points and throughout the Maldives so that manta based tourism activities can be developed more successfully and the impact of tourism can be monitored. Using natural markings to identify individual manta rays, Petersen and Jolly-Seber probabilistic estimators were used to assess population size for main dive sites and the central atolls of North Male, Ari and Baa. Between 2000 and 2009, 1835 individual mantas were identified from 3373 sightings. There was a significant gender bias with a male: female ratio of 1:1.8. Only 24.7% of manta rays were re-sighted between 1 and 34 times with a maximum gap of 7 years and 11 months between sightings. Estimates for populations around main dive sites were 181–562. The estimated population of M. alfredi for North Male atoll is 537 (SE ± 53.2). Estimated atoll populations were extrapolated to obtain an estimate of Manta alfredi population for the entire Maldives of 9,677 individuals.     

Manta rays in the Marquesas Islands: first records of Manta birostris in French Polynesia and most easterly location of Manta alfredi in the Pacific Ocean,with notes on their distribution

Based on direct observations of free‐ranging specimens, the giant manta ray Manta birostris is reported from the Marquesas Islands, the first sighting in French Polynesia. Sightings of its sister species, the reef manta ray Manta alfredi, are also reported at the most easterly location in the Pacific Ocean. Preliminary individual identification as well as notes on their distribution are also reported.     

Observations of mating behavior of the manta ray,manta birostris, at the ogasawara islands, japan

On 11 July 1997, the mating behavior of wild manta rays,Manta birostris, was observed while skin diving off Chichijima. Ogasawara Islands, Japan, and recorded with 49 underwater photographs and about 20 minutes of video tape. The female manta ray involved was estimated as being approximately 5 m in dise width (DW) and the two males involved, approximately 4 m DW. Copulatory behavior of the two males appeared to be almost the same, copulation itself being of the abdomen-to-abdomen type. Initially, the males chased the female for 20–30 minutes, all animals swimming at approximately 10 km/h. Each copulation event occurred within one meter of the surface, during which time the participating male grasped the tip of the female's left pectoral fin with his mouth. The clasper was inserted for 90 seconds (Male No. 1) and 60 seconds (Male No. 2), respectively. The mating behavior sequence of the manta rays involved following five steps. 1.: Male chases behind the tail of the female, attempting (several times) to grasp the latter's pectoral fin (chasing behavior). 2.: Male bites the tip of the female's pectoral fin, before positioning itself against the latter's underside (biting behavior). 3.: Male inserts a clasper into the cloaca of the female (copulating behavior). 4.: Male removes the clasper from the cloaca of the female, but continues biting the latter's pectoral fin (post-copulating behavior). 5.: Male releases the pectoral fin of the female, setting her free (separating behavior).     

Sex biased individual variation in movement patterns of a highly mobile,near-shore marine planktivore,the reef manta ray Mobula alfredi

We examined individual variation and the role of sex on the movements of the reef manta ray Mobula alfredi. Specifically, we analysed several movement metrics using 6 years of nightly observations (1 January 2009–31 December 2014) of 118 individually identifiable manta rays at two discrete but spatially proximate sites, locally known as Manta Heaven and Manta Village, 15 km apart on the west side of the island of Hawaii, USA. Males were slightly more often (33.5%, model fitted mean, P < 0.05) observed than females at Manta Heaven, but females were much more often (156.4%, model fitted mean, P < 0.05) observed at Manta Village. Movement patterns among individuals varied greatly, but the level of variation was similar between sexes. Some animals, mainly females, displayed more resident patterns, whereas other, more mobile, animals moved between sites more frequently and had longer gaps between sightings. We did not detect discrete behavioural groups; rather, individuals varied along a continuous spectrum from many observations and high affinity to few observations and low fidelity to survey locations. These complex and variable movement patterns observed at the individual level, between sexes and between two nearby sites, in Hawaii's manta rays highlight the need for finer scale considerations in conservation and management of highly mobile marine populations.     

Analysis and Human Health Risk Assessment of Arsenic,Cadmium, and Mercury in Manta Birostris (Manta Ray) Caught Along the Ghanaian Coastline

The Ghanaian marine water of the Atlantic Ocean could be said to be polluted with As and Hg. However, Cd levels do not appear to pose an environmental concern based on levels reported in tissue samples collected. Tissue samples from six Manta birostris (manta ray) used as a bioindicator were tested for these heavy metals because many people in Ghana eat this fish (aka “kako”). The pattern of bioaccumulation of these heavy metals found in the tissues of manta rays, which feed on phytoplankton, suggests that pollution has traveled deeper into the sea, which is an environmental concern and one that may require some urgent attention. Concentrations of As, Cd, and Hg were determined in kidney, liver, and muscle samples from six Manta birostris using the neutron activation analysis method. The results showed elevated levels of the metals in the manta rays. If these values are interpreted to represent daily human exposure estimates of possible human health cancer and non-cancer risks may be derived to conclude some potential risk for daily consumers of manta ray tissues. From the assessment, most of the hazard indexes for children were less than 1 except for arsenic, which was greater than 1. This means that there is for arsenic some hazard risk for children consumers of the manta ray. This is the first study to show the accumulation of heavy metals in manta ray; although the sources are not very clear, and may be due to the numerous mining activities, it needs to be investigated further. The presence of the heavy metals is very significant because the meat (muscle) is consumed in Ghana and hence there is possible accumulation in humans.     

Reproductive ecology of the reef manta ray Manta alfredi in southern Mozambique

The application of a photographic identification methodology using the unique ventral surface markings (natural spot patterns) of an observed population in southern Mozambique enabled many aspects of the reproductive ecology of reef manta rays Manta alfredi to be examined. The region encompassing the study site was identified as a mating ground for M. alfredi based on observations of mating events and fresh mating scars on females. The distribution of these pectoral fin scars was highly biased and indicated a strong lateralized behavioural trait, with 99% of these scars occurring only on the left pectoral fin. No other elasmobranch has been reported to display behavioural lateralization. The study region also acts as a birthing ground, with individuals typically giving birth in the austral summer period after a gestation of c. 1 year. Reproductive periodicity in M. alfredi was most commonly biennial, but a few individuals were pregnant in consecutive years, confirming an annual ovulatory cycle. The production of a single pup appears to be the normal situation, although observations in the wild as well as during opportunistic dissections of individuals killed by fisheries revealed that two pups are conceived on occasion. Many aspects of the study have contributed to the limited baseline data currently available for this species and have highlighted the potential need for more conservative conservation strategies.     

Satellite tracking of manta rays highlights challenges to their conservation

We describe the real-time movements of the last of the marine mega-vertebrate taxa to be satellite tracked - the giant manta ray (or devil fish, Manta birostris), the world's largest ray at over 6 m disc width. Almost nothing is known about manta ray movements and their environmental preferences, making them one of the least understood of the marine mega-vertebrates. Red listed by the International Union for the Conservation of Nature as 'Vulnerable' to extinction, manta rays are known to be subject to direct and incidental capture and some populations are declining. Satellite-tracked manta rays associated with seasonal upwelling events and thermal fronts off the Yucatan peninsula, Mexico, and made short-range shuttling movements, foraging along and between them. The majority of locations were received from waters shallower than 50 m deep, representing thermally dynamic and productive waters. Manta rays remained in the Mexican Exclusive Economic Zone for the duration of tracking but only 12% of tracking locations were received from within Marine Protected Areas (MPAs). Our results on the spatio-temporal distribution of these enigmatic rays highlight opportunities and challenges to management efforts.     

Distribution of the reef manta ray Mobula alfredi and the oceanic manta ray Mobula birostris in the Philippines: a collaborative effort for conservation

Little is known about manta ray population size, structure and connectivity in the Philippines. In collaboration with dive operators, non-governmental organizations and authorities, sightings of manta rays were collated into a single national database. Using in-water photographs and videos gathered through citizen science and dedicated research efforts, this study compiled sightings between 2004 and 2020, showing 22 separate sites throughout the archipelago with manta rays present. A total of 392 individual reef manta rays (Mobula alfredi) and 107 oceanic manta rays (Mobula birostris) were identified from the collected footage. Four specific sites in the provinces of Masbate and Palawan together hosted 89% of all identified individuals and accounted for 95% of sightings, highlighting these areas are key aggregation sites. This study also reports the movements of M. birostris within the Philippines, based on photo-identification of three individuals moving 150 km between Cebu and Masbate. Despite the growing number of recreational divers in Daanbantayan and San Jacinto, an 80% decline in M. birostris sightings was observed at these sites. To ensure effective future conservation, it is recommended that efforts focus on the identification and protection of manta ray hotspots and migratory corridors, the creation of a sustainable tourism framework and, most important, the implementation of mitigation strategies to reduce fisheries interactions.     

Notes on the feeding behaviour and swimming pattern of Manta alfredi (Chondrichthyes,Mobulidae) in the Red Sea

The feeding behaviour and the swimming pattern of the reef manta (Manta alfredi) are described based on video recordings conducted in Sudan, Red Sea, Indian Ocean, during June 2001. Three circular swimming patterns are described: horizontal circle pattern, inclined circle pattern and double circle pattern, related to feedings on zooplankton distributed in the water column. The observation of circle-like swimming patterns for reef manta rays in shallow waters expands our knowledge about M. alfredi and suggests that the distribution and abundance of zooplankton influence the occurrence and group size of this species in the studied area, as reported from other localities for Manta species.     

Occurrence and use of an estuarine habitat by giant manta ray Manta birostris

Based on the knowledge of local artisanal fishermen and on direct observations, this study presents evidence that the giant manta ray Manta birostris uses the Paranaguá estuarine complex in south Brazil, south‐western Atlantic Ocean, in a predictable seasonal pattern. Behavioural observations suggest that the estuary can act as a nursery ground for M. birostris during the summer.     

Contingency checking and self-directed behaviors in giant manta rays: Do elasmobranchs have self-awareness?

Elaborate cognitive skills arose independently in different taxonomic groups. Self-recognition is conventionally identified by the understanding that one’s own mirror reflection does not represent another individual but oneself, which has never been proven in any elasmobranch species to date. Manta rays have a high encephalization quotient, similar to those species that have passed the mirror self-recognition test, and possess the largest brain of all fish species. In this study, mirror exposure experiments were conducted on two captive giant manta rays to document their response to their mirror image. The manta rays did not show signs of social interaction with their mirror image. However, frequent unusual and repetitive movements in front of the mirror suggested contingency checking; in addition, unusual self-directed behaviors could be identified when the manta rays were exposed to the mirror. The present study shows evidence for behavioral responses to a mirror that are prerequisite of self-awareness and which has been used to confirm self-recognition in apes.     

Courtship and mating behaviour of manta rays Mobula alfredi and M. birostris in the Maldives

The aim of this 14 year study was to elucidate the entire courtship and mating behaviour of manta rays Mobula alfredi and M. birostris using behavioural observations, video and photographic records. From 2003 to 2016, over 11,000 surveys were undertaken at known manta ray aggregation sites in the Maldives to record any observed manta rays reproductive activity. From 47,591 photo‐ID sightings, 4,247 individual M. alfredi were identified and 226 individual M. birostris from 229 photo‐ID sightings, all recorded at 22 atolls across 265 different sites. Courtship activity was observed on 206 surveys at 30 different sites. A total of 229 courtship events were recorded, with 90% (n = 205) of them occurring at cleaning sites. The observed courtship activity was categorised into seven distinct stages which are described in detail: initiation, endurance, evasion, pre‐copulatory positioning, copulation, post‐copulatory holding and separation. Photographs provide the first scientific record of the entirety of manta rays courtship and mating. Both M. alfredi and M. birostris appear to engage in the same elaborate courtship rituals, exhibiting the same behaviours during all stages of the courtship and mating process.     

How common is common? Rapidly assessing population size and structure of the frog Mantidactylus betsileanus at a site in east‐central Madagascar

Population‐level data are urgently needed for amphibians in light of the ongoing amphibian extinction crisis. Studies focused on population dynamics are not only important for rare species but also for common species which shape ecosystems to a greater degree than those that are rare. Some of the greatest global amphibian species diversity is found in Madagascar, yet there are few studies on the ecology of frog species on the island. We carried out a mark‐recapture study on the widespread frog Mantidactylus betsileanus (Mantellidae: Mantellinae: Mantidactylus) at two adjacent rainforest sites in east‐central Madagascar to assess its population size and structure. To do so, we validated and implemented an individual identification protocol using photographs of the ventral patterns of frogs and identified individuals with photographic‐matching software. Using this rapid, non‐invasive survey method, we were able to estimate a density of 26 and 28 frogs per 100 m² at each of the two sites sampled. Our results show the rainforests near the village of Andasibe, Madagascar support remarkably high amphibian abundance, helping illustrate the significant ecological role of frogs in this ecosystem. Further, individual frog markings allowed us to develop more precise estimates than traditional survey methods. This study provides a blueprint to augment existing population studies or develop new monitoring programs in Madagascar and beyond.     

The reproductive ecology of resident manta rays (Manta alfredi) off Maui, Hawaii, with an emphasis on body size

In resident manta rays (Manta alfredi) off Maui, sexual maturity appears delayed until growth exceeds 90% of maximum size, an indicator that large body size provides a reproductive advantage at the expense of a shorter reproductive time period. In this study, 286 surveys were conducted between 2005 and 2010 using photo-identification and photogrammetry to study the reproductive ecology of a resident population of manta rays off Maui, Hawaii, and investigate the reproductive benefits of large body size in each sex. Although reproductive activities occurred year-round, mating trains and late-term pregnant females were significantly more likely to be observed during the winter months. Some females were pursued by males during both winter and summer of the same year, suggesting multiple ovulations may be possible in a single year. Males likely detect a female’s reproductive state by positioning directly behind her, or passing through her bodily excretions. The mean pregnancy rate was estimated at 0.56 pregnancies/adult female/year with larger females pregnant more often, and more likely in consecutive years. The operational sex ratio was heavily skewed with 2.68 adult males per reproductively available female. Although males appear to compete with one another for females within a mating train, no direct physical competition was ever observed between males. Evidence of highly dynamic mating trains lasting more than one day suggests endurance rivalry may be the primary mating strategy among males, during which larger males may benefit from greater energy reserves. The study area appears to be an important staging area for mating individuals in this population.     

Genetic fingerprinting proves cross‐correlated automatic photo‐identification of individuals as highly efficient in large capture–mark–recapture studies

Capture–mark–recapture (CMR) approaches are the backbone of many studies in population ecology to gain insight on the life cycle, migration, habitat use, and demography of target species. The reliable and repeatable recognition of an individual throughout its lifetime is the basic requirement of a CMR study. Although invasive techniques are available to mark individuals permanently, noninvasive methods for individual recognition mainly rest on photographic identification of external body markings, which are unique at the individual level. The re‐identification of an individual based on comparing shape patterns of photographs by eye is commonly used. Automated processes for photographic re‐identification have been recently established, but their performance in large datasets (i.e., > 1000 individuals) has rarely been tested thoroughly. Here, we evaluated the performance of the program AMPHIDENT, an automatic algorithm to identify individuals on the basis of ventral spot patterns in the great crested newt (Triturus cristatus) versus the genotypic fingerprint of individuals based on highly polymorphic microsatellite loci using GENECAP. Between 2008 and 2010, we captured, sampled and photographed adult newts and calculated for 1648 samples/photographs recapture rates for both approaches. Recapture rates differed slightly with 8.34% for GENECAP and 9.83% for AMPHIDENT. With an estimated rate of 2% false rejections (FRR) and 0.00% false acceptances (FAR), AMPHIDENT proved to be a highly reliable algorithm for CMR studies of large datasets. We conclude that the application of automatic recognition software of individual photographs can be a rather powerful and reliable tool in noninvasive CMR studies for a large number of individuals. Because the cross‐correlation of standardized shape patterns is generally applicable to any pattern that provides enough information, this algorithm is capable of becoming a single application with broad use in CMR studies for many species.     

Photographic identification of individuals of a free‐ranging,small terrestrial vertebrate

Recognition of individuals within an animal population is central to a range of estimates about population structure and dynamics. However, traditional methods of distinguishing individuals, by some form of physical marking, often rely on capture and handling which may affect aspects of normal behavior. Photographic identification has been used as a less‐invasive alternative, but limitations in both manual and computer‐automated recognition of individuals are particularly problematic for smaller taxa (<500 g). In this study, we explored the use of photographic identification for individuals of a free‐ranging, small terrestrial reptile using (a) independent observers, and (b) automated matching with the Interactive Individual Identification System (I³S Pattern) computer algorithm. We tested the technique on individuals of an Australian skink in the Egernia group, Slater's skink Liopholis slateri, whose natural history and varied scale markings make it a potentially suitable candidate for photo‐identification. From ‘photographic captures’ of skink head profiles, we designed a multi‐choice key based on alternate character states and tested the abilities of observers — with or without experience in wildlife survey — to identify individuals using categorized test photos. We also used the I³S Pattern algorithm to match the same set of test photos against a database of 30 individuals. Experienced observers identified a significantly higher proportion of photos correctly (74%) than those with no experience (63%) while the I³S software correctly matched 67% as the first ranked match and 83% of images in the top five ranks. This study is one of the first to investigate photo identification with a free‐ranging small vertebrate. The method demonstrated here has the potential to be applied to the developing field of camera‐traps for wildlife survey and thus a wide range of survey and monitoring applications.     

Empirical tests of the role of disruptive coloration in reducing detectability

Disruptive patterning is a potentially universal camouflage technique that is thought to enhance concealment by rendering the detection of body shapes more difficult. In a recent series of field experiments, artificial moths with markings that extended to the edges of their 'wings' survived at higher rates than moths with the same edge patterns inwardly displaced. While this result seemingly indicates a benefit to obscuring edges, it is possible that the higher density markings of the inwardly displaced patterns concomitantly reduced their extent of background matching. Likewise, it has been suggested that the mealworm baits placed on the artificial moths could have created differential contrasts with different moth patterns. To address these concerns, we conducted controlled trials in which human subjects searched for computer-generated moth images presented against images of oak trees. Moths with edge-extended disruptive markings survived at higher rates, and took longer to find, than all other moth types, whether presented sequentially or simultaneously. However, moths with no edge markings and reduced interior pattern density survived better than their high-density counterparts, indicating that background matching may have played a so-far unrecognized role in the earlier experiments. Our disruptively patterned non-background-matching moths also had the lowest overall survivorship, indicating that disruptive coloration alone may not provide significant protection from predators. Collectively, our results provide independent support for the survival value of disruptive markings and demonstrate that there are common features in human and avian perception of camouflage.